leopard

The English name "leopard" comes from Old French {{lang|fro|leupart}} or Middle French {{lang|frm|liepart}}, that derives from Latin {{lang|la|leopardus}} and ancient Greek {{lang|grc|λέοπάρδος}} ({{transl|grc|leopardos}}). {{transl|grc|Leopardos}} could be a compound of {{lang|grc|λέων}} ({{transl|grc|leōn}}), meaning {{gloss|lion}}, and {{lang|grc|πάρδος}} ({{transl|grc|pardos}}), meaning {{gloss|spotted}}.{{cite book |last1=Lewis |first1=C. T. |author1-link=Charlton Thomas Lewis |last2=Short |first2=C. |name-list-style=amp |year=1879 |title=A Latin Dictionary |location=Oxford |publisher=Clarendon Press |chapter=lěǒpardus |page=1069 |chapter-url=https://archive.org/details/in.ernet.dli.2015.61236/page/n1069}}{{cite book |last1=Liddell |first1=H. G. |author1-link=Henry Liddell |last2=Scott |first2=R. |author2-link=Robert Scott (philologist) |name-list-style=amp |year=1889 |title=A Greek–English Lexicon |location=Oxford |publisher=Clarendon Press |chapter=λέο-πάρδος |page=884 |chapter-url=https://archive.org/details/greekenglishlex00liddrich/page/884}}{{Cite book |last=Partridge |first=E. |author-link=Eric Partridge |title=Origins: A Short Etymological Dictionary of Modern English |publisher=Greenwich House |year=1983 |location=New York |isbn=978-0-517-41425-5 |page=349 |url=https://archive.org/details/originsshortetym0000part/page/349}} The word {{lang|drc|λέοπάρδος}} originally referred to a cheetah (Acinonyx jubatus).{{Cite journal |last=Nicholas |first=N. |s2cid=56160515 |year=1999|title=A conundrum of cats: pards and their relatives in Byzantium |journal=Greek, Roman, and Byzantine Studies |volume=40 |pages=253–298}}

"Panther" is another common name, derived from Latin {{lang|la|panther}} and ancient Greek {{lang|grc|πάνθηρ}} ({{transl|grc|pánthēr}}); The generic name Panthera originates in Latin {{lang|la|panthera}}, a hunting net for catching wild beasts to be used by the Romans in combats.{{cite book |last1=Lewis |first1=C. T. |last2=Short |first2=C. |name-list-style=amp |year=1879 |title=A Latin Dictionary |location=Oxford |publisher=Clarendon Press |chapter=panthera |page=1298 |chapter-url=https://archive.org/details/in.ernet.dli.2015.61236/page/n1317}} {{lang|la|Pardus}} is the masculine singular form.{{cite book |last1=Lewis |first1=C. T. |last2=Short |first2=C. |name-list-style=amp |year=1879 |title=A Latin Dictionary |location=Oxford |publisher=Clarendon Press |chapter=pardus |pages=1302 |chapter-url=https://archive.org/details/in.ernet.dli.2015.61236/page/n1321}}

File:Panthera pardus subspecies map.png

Felis pardus was the scientific name proposed by Carl Linnaeus in 1758.{{cite book |author=Linnaeus, C. |year=1758 |title=Caroli Linnæi Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis |volume=Tomus I |edition=decima, reformata |location=Holmiae |publisher=Laurentius Salvius |pages=41−42 |chapter=Felis pardus |chapter-url=https://archive.org/stream/mobot31753000798865#page/41/mode/2up}} {{in lang|la}}

The generic name Panthera was first used by Lorenz Oken in 1816, who included all the known spotted cats into this group.{{cite book |last1=Oken |first1=L. |year=1816 |title=Lehrbuch der Zoologie. 2. Abtheilung |location=Jena |publisher=August Schmid & Comp. |page=1052 |chapter=1. Art, Panthera |chapter-url=https://books.google.com/books?id=S5o5AAAAcAAJ&pg=PA1052 |access-date=2019-04-06 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145524/https://books.google.com/books?id=S5o5AAAAcAAJ&pg=PA1052#v=onepage&q&f=false |url-status=live }}

Oken's classification was not widely accepted, and Felis or Leopardus was used as the generic name until the early 20th century.{{cite book|last1=Ellerman |first1=J. R. |last2=Morrison-Scott |first2=T. C. S. |year=1966 |url=https://archive.org/stream/checklistofindia00elle#page/314/mode/2up |title=Checklist of Palaearctic and Indian mammals 1758 to 1946 |edition=Second|publisher=British Museum of Natural History |location= London |pages=315–317}}

The leopard was designated as the type species of Panthera by Joel Asaph Allen in 1902.{{cite journal |last1=Allen |first1=J. A. |year=1902 |title=Mammal names proposed by Oken in his 'Lehrbuch der Zoologie' |journal=Bulletin of the American Museum of Natural History |volume=16 |issue=27 |pages=373−379 |url=http://digitallibrary.amnh.org/items/bc682fbb-d0d7-42a1-9906-bb50b3b0ae31 |access-date=2024-03-14 |archive-date=2023-12-09 |archive-url=https://web.archive.org/web/20231209013613/https://digitallibrary.amnh.org/items/bc682fbb-d0d7-42a1-9906-bb50b3b0ae31 |url-status=live }}

In 1917, Reginald Innes Pocock also subordinated the tiger (P. tigris), lion (P. leo), and jaguar (P. onca) to Panthera.{{cite journal |last1=Pocock |first1=R. I. |year=1917 |title=The Classification of existing Felidae |journal=The Annals and Magazine of Natural History |series=Series 8 |volume=XX |pages=329–350 |doi=10.1080/00222931709487018 |url=https://archive.org/stream/annalsmagazineof8201917lond#page/n359/mode/2up}}{{cite book |last=Pocock |first= R. I. |year=1939 |title=The Fauna of British India, including Ceylon and Burma. Mammalia: Volume 1 |publisher=Taylor and Francis |location=London |pages=222–239 |chapter=Panthera pardus |chapter-url=https://archive.org/stream/PocockMammalia1/pocock1#page/n273/mode/2up}}

Following Linnaeus' first description, 27 leopard subspecies were proposed by naturalists between 1794 and 1956. Since 1996, only eight subspecies have been considered valid on the basis of mitochondrial analysis.{{cite journal |last1=Miththapala |first1=S. |last2=Seidensticker |first2=J. |last3=O'Brien |first3=S. J. |name-list-style=amp |title=Phylogeographic subspecies recognition in leopards (Panthera pardus): molecular genetic variation |journal=Conservation Biology |year=1996 |volume=10 |issue=4 |pages=1115–1132 |doi=10.1046/j.1523-1739.1996.10041115.x |bibcode=1996ConBi..10.1115M |url=https://repository.si.edu/bitstream/handle/10088/4298/Miththapala1996.pdf?sequence=1&isAllowed=y |access-date=2017-03-09 |archive-date=2016-02-17 |archive-url=https://web.archive.org/web/20160217074413/https://repository.si.edu/bitstream/handle/10088/4298/Miththapala1996.pdf?sequence=1&isAllowed=y |url-status=live }} Later analysis revealed a ninth valid subspecies, the Arabian leopard.{{cite journal |last1=Uphyrkina |first1=O. |last2=Johnson |first2=E. W. |last3=Quigley |first3=H. |last4=Miquelle |first4=D. |last5=Marker |first5=L. |last6=Bush |first6=M. |last7=O'Brien |first7=S. J. |name-list-style=amp |year=2001 |title=Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus |url=http://www.biosoil.ru/files/00001386.pdf |url-status=live |journal=Molecular Ecology |volume=10 |issue=11 |pages=2617–2633 |doi=10.1046/j.0962-1083.2001.01350.x |pmid=11883877 |bibcode=2001MolEc..10.2617U |s2cid=304770 |archive-url=https://web.archive.org/web/20110910162803/http://www.biosoil.ru/files/00001386.pdf |archive-date=2011-09-10}}

In 2017, the Cat Classification Task Force of the Cat Specialist Group recognized the following eight subspecies as valid taxa:{{cite journal |author1=Kitchener, A. C. |author2=Breitenmoser-Würsten, C. |author3=Eizirik, E. |author4=Gentry, A. |author5=Werdelin, L. |author6=Wilting, A. |author7=Yamaguchi, N. |author8=Abramov, A. V. |author9=Christiansen, P. |author10=Driscoll, C. |author11=Duckworth, J. W. |author12=Johnson, W. |author13=Luo, S.-J. |author14=Meijaard, E. |author15=O’Donoghue, P. |name-list-style=amp |author16=Sanderson, J. |author17=Seymour, K. |author18=Bruford, M. |author19=Groves, C. |author20=Hoffmann, M. |author21=Nowell, K. |author22=Timmons, Z. |author23=Tobe, S. |year=2017 |title=A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group |journal=Cat News |issue=Special Issue 11 |pages=73–75 |url=https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=73 |access-date=2019-07-25 |archive-date=2020-01-17 |archive-url=https://web.archive.org/web/20200117172708/https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=73 |url-status=live}}

Results of an analysis of molecular variance and pairwise fixation index of 182 African leopard museum specimens showed that some African leopards exhibit higher genetic differences than Asian leopard subspecies.{{cite journal |author1=Anco, C. |author2=Kolokotronis, S. O. |author3=Henschel, P. |author4=Cunningham, S. W. |author5=Amato, G. |author6=Hekkala, E. |name-list-style=amp |title=Historical mitochondrial diversity in African leopards (Panthera pardus) revealed by archival museum specimens |journal=Mitochondrial DNA Part A |volume=29 |issue=3 |pages=455–473 |doi=10.1080/24701394.2017.1307973 |pmid=28423965 |year=2017 |s2cid=4348541}}

File:Two cladograms for Panthera.svg

Results of phylogenetic studies based on nuclear DNA and mitochondrial DNA analysis showed that the last common ancestor of the Panthera and Neofelis genera is thought to have lived about {{Ma|6.37}}. Neofelis diverged about {{Ma|8.66}} from the Panthera lineage. The tiger diverged about {{Ma|6.55}}, followed by the snow leopard about {{Ma|4.63}} and the leopard about {{Ma|4.35}}. The leopard is a sister taxon to a clade within Panthera, consisting of the lion and the jaguar.{{cite journal |last1=Johnson |first1=W. E. |last2=Eizirik |first2=E. |last3=Pecon-Slattery |first3=J. |last4=Murphy |first4=W. J. |last5=Antunes |first5=A. |last6=Teeling |first6=E. |last7=O'Brien |first7=S. J. |title=The late Miocene radiation of modern Felidae: a genetic assessment |journal=Science |year=2006 |volume=311 |issue=5757 |pages=73–77 |name-list-style=amp |pmid=16400146 |doi=10.1126/science.1122277 |bibcode=2006Sci...311...73J |s2cid=41672825 |url=https://zenodo.org/record/1230866 |access-date=2019-07-05 |archive-date=2020-10-04 |archive-url=https://web.archive.org/web/20201004075725/https://zenodo.org/record/1230866 |url-status=live }}{{cite book |last1=Werdelin |first1=L. |last2=Yamaguchi |first2=N. |last3=Johnson |first3=W. E. |last4=O'Brien |first4=S. J. |name-list-style=amp |chapter=Phylogeny and evolution of cats (Felidae) |year=2010 |pages=59–82 |chapter-url=https://www.researchgate.net/publication/266755142 |editor1-last=Macdonald |editor1-first=D. W. |editor2-last=Loveridge |editor2-first=A. J. |title=Biology and Conservation of Wild Felids |publisher=Oxford University Press |location=Oxford, UK |isbn=978-0-19-923445-5 |access-date=2018-09-25 |archive-date=2018-09-25 |archive-url=https://web.archive.org/web/20180925141956/https://www.researchgate.net/publication/266755142 |url-status=live }}

Results of a phylogenetic analysis of chemical secretions amongst cats indicated that the leopard is closely related to the lion.{{cite journal |last1=Bininda-Emonds |first1=O. R. P. |last2=Decker-Flum |first2=D. M. |last3=Gittleman |first3=J. L. |name-list-style=amp |title=The utility of chemical signals as phylogenetic characters: an example from the Felidae |journal=Biological Journal of the Linnean Society |year=2001 |volume=72 |issue=1 |pages=1–15 |doi=10.1111/j.1095-8312.2001.tb01297.x |doi-access=free}}

The geographic origin of the Panthera is most likely northern Central Asia. The leopard-lion clade was distributed in the Asian and African Palearctic since at least the early Pliocene.{{cite journal |author1=Tseng, Z. J. |author2=Wang, X. |author3=Slater, G. J. |author4=Takeuchi, G. T. |author5=Li, Q. |author6=Liu, J. |author7=Xie, G. |name-list-style=amp |date=2014 |title=Himalayan fossils of the oldest known pantherine establish ancient origin of big cats |journal=Proceedings of the Royal Society B: Biological Sciences |volume=281 |issue=1774 |page=20132686 |doi=10.1098/rspb.2013.2686|pmid=24225466 |pmc=3843846 }} The leopard-lion clade diverged 3.1–1.95 million years ago.{{cite journal |last1=Davis |first1=B. W. |last2=Li |first2=G. |last3=Murphy |first3=W. J. |name-list-style=amp |title=Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, Panthera (Carnivora: Felidae) |journal=Molecular Phylogenetics and Evolution |year=2010 |volume=56 |issue=1 |pages=64–76 |doi=10.1016/j.ympev.2010.01.036 |pmid=20138224 |bibcode=2010MolPE..56...64D |url=http://web.csulb.edu/~acarter3/course-evolution/files/cat-phylogeny.pdf |url-status=dead |archive-url=https://web.archive.org/web/20160305131606/http://web.csulb.edu/~acarter3/course-evolution/files/cat-phylogeny.pdf |archive-date=2016-03-05}}{{cite journal |last1=Mazák |first1=J. H. |last2=Christiansen |first2=P. |last3=Kitchener|first3=A. C. |last4=Goswami |first4=A. |name-list-style=amp |title=Oldest known pantherine skull and evolution of the tiger |journal=PLOS ONE |year=2011 |volume=6 |issue=10 |pages=e25483 |doi=10.1371/journal.pone.0025483 |pmid=22016768 |pmc=3189913 |bibcode=2011PLoSO...625483M |doi-access=free}} Additionally, a 2016 study revealed that the mitochondrial genomes of the leopard, lion and snow leopard are more similar to each other than their nuclear genomes, indicating that their ancestors hybridized with the snow leopard at some point in their evolution.{{cite journal |last1=Li |first1=G. |last2=Davis |first2=B. W. |last3=Eizirik |first3=E. |last4=Murphy |first4=W. J. |name-list-style=amp |title=Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae) |journal=Genome Research |date=2016 |volume=26 |issue=1 |pages=1–11 |doi=10.1101/gr.186668.114 |pmid=26518481 |pmc=4691742}}

The oldest unambiguous fossils of the leopard are from Eastern Africa, dating to around 2 million years ago.{{Cite journal |last1=Paijmans |first1=J. L. A. |last2=Barlow |first2=A. |last3=Förster |first3=D. W. |last4=Henneberger |first4=K. |last5=Meyer |first5=M. |last6=Nickel |first6=B. |last7=Nagel |first7=D. |last8=Havmøller |first8=R. W. |last9=Baryshnikov |first9=G. F. |last10=Joger |first10=U. |last11=Rosendahl |first11=W. |last12=Hofreiter |first12=M. |name-list-style=amp |date=2018 |title=Historical biogeography of the leopard (Panthera pardus) and its extinct Eurasian populations |journal=BMC Evolutionary Biology |volume=18 |issue=1 |page=156 |doi=10.1186/s12862-018-1268-0 |pmc=6198532 |pmid=30348080 |doi-access=free |bibcode=2018BMCEE..18..156P}}

Leopard-like fossil bones and teeth possibly dating to the Pliocene were excavated in Perrier in France, northeast of London, and in Valdarno, Italy. Until 1940, similar fossils dating back to the Pleistocene were excavated mostly in loess and caves at 40 sites in Europe, including Furninha Cave near Lisbon, Genista Caves in Gibraltar, and Santander Province in northern Spain to several sites across France, Switzerland, Italy, Austria, Germany, in the north up to Derby in England, in the east to Přerov in the Czech Republic and the Baranya in southern Hungary.{{cite journal |last=Schmid |first= E. |year=1940 |title=Variationstatistische Untersuchungen am Gebiss pleistozäner und rezenter Leoparden und anderer Feliden |journal=Zeitschrift für Säugetierkunde |volume=15 |pages=1–179 |url=https://archive.org/details/zeitschrift1519401943deut/page/n11}}

Leopards arrived in Eurasia during the late Early to Middle Pleistocene around 1.2{{Cite journal |last1=Marciszak |first1=A. |last2=Lipecki |first2=G. |last3=Gornig |first3=W. |last4=Matyaszczyk |first4=L. |last5=Oszczepalińska |first5=O. |last6=Nowakowski |first6=D. |last7=Talamo |first7=S. |title=The first radiocarbon-dated remains of the Leopard Panthera pardus (Linnaeus, 1758) from the Pleistocene of Poland |date=2022 |journal=Radiocarbon |volume=64 |issue=6 |pages=1359–1372 |doi=10.1017/RDC.2022.33 |bibcode=2022Radcb..64.1359M |s2cid=255850069 |hdl=11585/887180 |hdl-access=free |url=https://www.cambridge.org/core/product/identifier/S0033822222000339/type/journal_article |access-date=2023-07-18 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145527/https://www.cambridge.org/core/journals/radiocarbon/article/first-radiocarbondated-remains-of-the-leopard-panthera-pardus-linnaeus-1758-from-the-pleistocene-of-poland/14B7D49EB6F90245554DCBA441123E3B |url-status=live }} to 0.6 million years ago.

Four European Pleistocene leopard subspecies were proposed. P. p. begoueni from the beginning of the Early Pleistocene was replaced about {{Ma|0.6}} by P. p. sickenbergi, which in turn was replaced by P. p. antiqua around 0.3 million years ago. P. p. spelaea is the most recent subspecies that appeared at the beginning of the Late Pleistocene and survived until about 11,000 years ago and possibly into the early Holocene in the Iberian Peninsula.{{cite journal |last1=Diedrich|first1=C. G. |title=Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art |journal=Quaternary Science Reviews |year=2013 |volume=76 |pages=167–193 |doi=10.1016/j.quascirev.2013.05.009 |bibcode=2013QSRv...76..167D}}{{Cite journal |last1=Sauqué |first1=V. |name-list-style=amp |last2=Cuenca‑Bescós |first2=G. |date=2013 |title=The Iberian Peninsula, the last European refugium of Panthera pardus Linnaeus 1758 during the Upper Pleistocene |journal=Quaternaire |url=https://journals.openedition.org/quaternaire/6468 |volume=24 |issue=1 |pages=13–24 |doi=10.4000/quaternaire.6468 |access-date=2023-07-18 |archive-date=2023-07-18 |archive-url=https://web.archive.org/web/20230718210435/https://journals.openedition.org/quaternaire/6468 |url-status=live }}

Leopards depicted in cave paintings in Chauvet Cave provide indirect evidence of leopard presence in Europe.

Leopard fossils dating to the Late Pleistocene were found in Biśnik Cave in south-central Poland.{{Cite journal |last1=Marciszak |first1=A. |last2=Stefaniak |first2=K. |name-list-style=amp |date=2010 |title=Two forms of cave lion: Middle Pleistocene Panthera spelaea fossilis Reichenau, 1906 and Upper Pleistocene Panthera spelaea spelaea Goldfuss, 1810 from the Bísnik Cave, Poland |url=https://www.researchgate.net/publication/233669138 |journal=Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen |volume=258 |issue=3 |pages=339–351 |doi=10.1127/0077-7749/2010/0117 |access-date=2017-08-24 |archive-date=2018-09-25 |archive-url=https://web.archive.org/web/20180925142142/https://www.researchgate.net/publication/233669138 |url-status=live }}

Fossil remains were also excavated in the Iberian{{Cite journal |last1=Sauqué |first1=V. |last2=Rabal-Garcés |first2=R. |last3=Cuenca-Bescós |first3=G. |name-list-style=amp |date=2016 |title=Carnivores from Los Rincones, a leopard den in the highest mountain of the Iberian range (Moncayo, Zaragoza, Spain) |journal=Historical Biology |volume=28 |issue=4 |pages=479–506 |doi=10.1080/08912963.2014.977882 |doi-access=free |bibcode=2016HBio...28..479S}}{{Cite journal |last1=Sanchis |first1=A. |last2=Real |first2=C. |last3=Sauqué |first3=V. |last4=Núñez-Lahuerta |first4=C. |last5=Égüez |first5=N. |last6=Tormo |first6=C. |last7=Ripoll |first7=M. P. |last8=Carrión |first8=Y. M. |last9=Duarte |first9=E. |last10=de la Rasilla |first10=M. |name-list-style=amp |date=2019 |title=Neanderthal and carnivore activities at Llonin Cave, Asturias, northern Iberian Peninsula: Faunal study of Mousterian levels (MIS 3) |journal=Comptes Rendus Palevol |volume=18 |issue=1 |pages=113–141 |bibcode=2019CRPal..18..113S |doi=10.1016/j.crpv.2018.06.001 |s2cid=134157124 |hdl-access=free |hdl=10651/49880}} and Italian Peninsula,{{Cite journal |last1=Ghezzo |first1=E. |last2=Rook |first2=L. |date=2015 |name-list-style=amp |title=The remarkable Panthera pardus (Felidae, Mammalia) record from Equi (Massa, Italy): taphonomy, morphology, and paleoecology |journal=Quaternary Science Reviews |volume=110 |pages=131–151 |doi=10.1016/j.quascirev.2014.12.020}}{{Cite journal |last=Mecozzi |first=Beniamino |last2=Sardella |first2=Raffaele |last3=Boscaini |first3=Alberto |last4=Cherin |first4=Marco |last5=Costeur |first5=Loïc |last6=Madurell-Malapeira |first6=Joan |last7=Pavia |first7=Marco |last8=Profico |first8=Antonio |last9=Iurino |first9=Dawid A. |date=15 June 2021 |title=The tale of a short-tailed cat: New outstanding Late Pleistocene fossils of Lynx pardinus from southern Italy |url=https://www.sciencedirect.com/science/article/pii/S0277379121000470 |journal=Quaternary Science Reviews |language=en |volume=262 |pages=106840 |doi=10.1016/j.quascirev.2021.106840 |access-date=7 November 2024 |via=Elsevier Science Direct|hdl=2434/959858 |hdl-access=free }} and in the Balkans.{{Cite journal |last1=Sommer |first1=R. S. |last2=Benecke |first2=N. |name-list-style=amp |date=2006 |title=Late Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: a review |journal=Journal of Zoology |volume=269 |issue=1 |pages=7–19 |doi=10.1111/j.1469-7998.2005.00040.x}}{{Cite journal |last1=Miracle |first1=P. T. |last2=Lenardić |first2=J. M. |last3=Brajković |first3=D. |date=2010 |title=Last glacial climates, "Refugia", and faunal change in Southeastern Europe: Mammalian assemblages from Veternica, Velika pećina, and Vindija caves (Croatia) |journal=Quaternary International |volume=212 |issue=2 |pages=137–148 |doi=10.1016/j.quaint.2009.06.003|bibcode=2010QuInt.212..137M }}

Leopard fossils dating to the Pleistocene were also excavated in the Japanese archipelago.{{cite book |author=Izawa, M. |author2=Nakanishi, N. |name-list-style=amp |chapter=Felidae |pages=226−231 |editor=Ohdachi, S. D. |editor2=Ishibashi, Y. |editor3=Iwasa, M. A. |editor4=Saitoh, T. |year=2015 |title=The Wild Mammals of Japan |publisher=Shoukadoh Book Sellers and the Mammalogical Society of Japan |edition=Second |isbn=978-4-87974-691-7 |location=Kyoto}} Leopard fossils were also found in Taiwan.{{cite journal |author1=Chi T.-C. |name-list-style=amp |author2=Gan Y. |author3=Yang T.-R. |author4=Chang, C.-H. |year=2021 |title=First report of leopard fossils from a limestone cave in Kenting area, southern Taiwan |journal=PeerJ |volume=9 |page=e12020 |doi=10.7717/peerj.12020 |pmid=34513335 |pmc=8388558 |doi-access=free}}

{{Main|Panthera hybrid|Pumapard}}

In 1953, a male leopard and a female lion were crossbred in Hanshin Park in Nishinomiya, Japan. Their offspring known as a leopon was born in 1959 and 1961, all cubs were spotted and bigger than a juvenile leopard. Attempts to mate a leopon with a tigress proved unsuccessful.{{cite book |editor1=Kisling, V.N. |title=Zoo and Aquarium History : Ancient Animal Collections to Zoological Gardens |year=2001 |publisher=CRC Press |location=Boca Raton, Florida |isbn=978-0-8493-2100-9 |author=Kawata, K. |chapter=Zoological gardens of Japan |pages=295–329 |chapter-url=https://books.google.com/books?id=ulbMBQAAQBAJ&pg=PA314 |access-date=2021-09-07 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145322/https://books.google.com/books?id=ulbMBQAAQBAJ&pg=PA314#v=onepage&q&f=false |url-status=live }}

{{Multiple image |direction=vertical |align=right|image1=Leopard skeleton (black background).jpg|caption1=Mounted skeleton |image2=Panthera pardus 3d scan Natural History Museum University of Pisa C 1389.stl|caption2=3D model of skeleton|image3=Persian Leopard Fur 02.JPG |caption3=Rosettes of a leopard|image4=Blackleopard.JPG|caption4=A melanistic leopard or black panther}}

The leopard's fur is generally soft and thick, notably softer on the belly than on the back.{{cite book |year=2005 |title=The mammals of the southern African sub region |location=Cambridge |publisher=Cambridge University Press |isbn=9780521844185 |editor1-last=Skinner |editor1-first=J. D. |editor2-last=Chimimba |editor2-first=C. T. |edition=Third |last1=Mills |first1=M. G. L. |chapter=Subfamily Pantherinae |pages=385–396 |chapter-url=https://books.google.com/books?id=iqwEYkTDZf4C&pg=PA385 |access-date=2021-05-19 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505150034/https://books.google.com/books?id=iqwEYkTDZf4C&pg=PA385#v=onepage&q&f=false |url-status=live}} Its skin colour varies between individuals from pale yellowish to dark golden with dark spots grouped in rosettes. Its underbelly is white and its ringed tail is shorter than its body. Its pupils are round.{{cite book |last1=Mivart |first1=St. G. J. |author-link=St. George Jackson Mivart |title=The Cat: An Introduction to the Study of Backboned Animals, Especially Mammals |date=1900 |publisher=John Murray |location=London |chapter=Different kind of Cats |pages=391–439 |chapter-url=https://archive.org/details/catintroductiont00miva/page/394}} Leopards living in arid regions are pale cream, yellowish to ochraceous and rufous in colour; those living in forests and mountains are much darker and deep golden. Spots fade toward the white underbelly and the insides and lower parts of the legs.{{cite journal |last1=Pocook |first1=R. I. |year=1932 |title=The Leopards of Africa |journal=Proceedings of the Zoological Society of London |volume=102 |issue=2 |pages=543–591 |doi=10.1111/j.1096-3642.1932.tb01085.x}} Rosettes are circular in East African leopard populations, and tend to be squarish in Southern African and larger in Asian leopard populations. The fur tends to be grayish in colder climates, and dark golden in rainforest habitats.{{cite book |author=Nowell, K. |author2=Jackson, P. |name-list-style=amp |year=1996 |title=Wild Cats: status survey and conservation action plan |publisher=IUCN/SSC Cat Specialist Group |location=Gland, Switzerland |chapter=Leopard Panthera pardus (Linnaeus, 1758) |archive-url=https://web.archive.org/web/20140222223654/http://lynx.uio.no/lynx/catsgportal/cat-website/catfolk/ssaprd01.htm |archive-date=2014-02-22 |chapter-url=http://lynx.uio.no/lynx/catsgportal/cat-website/catfolk/ssaprd01.htm}} Rosette patterns are unique in each individual.{{cite book |last1=Schütze |first1=H. |title=Field Guide to the Mammals of the Kruger National Park |date=2002 |publisher=Struik Publishers|location=Cape Town, South Africa |isbn=978-1-86872-594-6 |pages=92–93}}{{cite book|last1=Menon |first1=V. |title=Indian Mammals: A Field Guide |date=2014 |publisher=Hachette |location=Gurgaon, India |isbn=978-93-5009-761-8}} This pattern is thought to be an adaptation to dense vegetation with patchy shadows, where it serves as camouflage.{{cite journal |author1=Allen, W. L. |author2=Cuthill, I. C. |author3=Scott-Samuel, N. E. |author4=Baddeley, R. |year=2010 |name-list-style=amp |title=Why the leopard got its spots: relating pattern development to ecology in felids |journal=Proceedings of the Royal Society B |volume=278 |issue=1710 |pages=1373–1380 |doi=10.1098/rspb.2010.1734 |pmc=3061134 |pmid=20961899}}

Its white-tipped tail is about {{cvt|60|-|100|cm|1}} long, white underneath and with spots that form incomplete bands toward the end of the tail.{{cite book |last1=Hoath |first1=R. |title=Field Guide to the Mammals of Egypt |date=2009 |publisher=American University in Cairo Press |location=Cairo, Egypt |isbn=978-977-416-254-1 |pages=106–107 |chapter=Leopard Panthera pardus (Linnaeus, 1758) |chapter-url=https://books.google.com/books?id=agWfg6oEKKkC&pg=PA106 |access-date=2021-05-19 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145927/https://books.google.com/books?id=agWfg6oEKKkC&pg=PA106#v=onepage&q&f=false |url-status=live}}

The guard hairs protecting the basal hairs are short, {{cvt|3-4|mm|1}} in face and head, and increase in length toward the flanks and the belly to about {{cvt|25|-|30|mm|1}}. Juveniles have woolly fur that appear to be dark-coloured due to the densely arranged spots.{{cite book |last=Estes |first=R. |year=1991 |title=The Behavior Guide to African Mammals, Including Hoofed Mammals, Carnivores, Primates |location=Los Angeles |publisher=The University of California Press |isbn=978-0-520-08085-0 |chapter=Leopard Panthera pardus |pages=366–369 |chapter-url=https://books.google.com/books?id=g977LsZHpcsC&pg=PA366 |access-date=2021-05-19 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145853/https://books.google.com/books?id=g977LsZHpcsC&pg=PA366#v=onepage&q&f=false |url-status=live}}

Its fur tends to grow longer in colder climates.{{cite journal |author1=Stein, A. B. |name-list-style=amp |author2=Hayssen, V. |title=Panthera pardus (Carnivora: Felidae) |journal=Mammalian Species |date=2010 |volume=45 |issue=900 |pages=30–48 |doi=10.1644/900.1 |s2cid=44839740 |doi-access=free|pmc=7149468 }}

The leopard's rosettes differ from those of the jaguar, which are darker and with smaller spots inside. The leopard has a diploid chromosome number of 38.{{Cite book |last1=Heptner |first1=V. G. |name-list-style=amp |last2=Sludskii |first2=A. A. |orig-year=1972 |year=1992 |title=Mlekopitajuščie Sovetskogo Soiuza. Moskva: Vysšaia Škola |trans-title=Mammals of the Soviet Union, Volume II, Part 2 |publisher=Smithsonian Institution and the National Science Foundation |location=Washington DC |chapter=Bars (leopard) |chapter-url=https://archive.org/stream/mammalsofsov221992gept#page/82/mode/2up |pages=203–273 |isbn=978-90-04-08876-4}}

Melanistic leopards are also known as black panthers. Melanism in leopards is caused by a recessive allele and is inherited as a recessive trait.{{cite journal|last1=Robinson |first1=R. |title=Inheritance of the black form of the leopard Panthera pardus |journal=Genetica |year=1970 |volume=41 |issue=1|pages=190–197 |doi=10.1007/BF00958904 |pmid=5480762 |s2cid=5446868}}{{cite journal |last1=Eizirik |first1=E. |last2=Yuhki |first2=N. |last3=Johnson |first3=W. E.|last4=Menotti-Raymond |first4=M. |last5=Hannah |first5=S. S. |last6=O'Brien |first6=S. J. |title=Molecular genetics and evolution of melanism in the cat family |journal=Current Biology |year=2003 |volume=13 |issue=5 |pages=448–453 |doi=10.1016/S0960-9822(03)00128-3 |pmid=12620197 |s2cid=19021807 |doi-access=free |bibcode=2003CBio...13..448E}}{{cite journal |last1=Kawanishi |first1=K. |last2=Sunquist |first2=M. E. |last3=Eizirik |first3=E. |last4=Lynam |first4=A. J. |last5=Ngoprasert |first5=D. |last6=Wan Shahruddin |first6=W. N. |last7=Rayan |first7=D. M. |last8=Sharma |first8=D. S. K.|last9=Steinmetz|first9=R. |name-list-style=amp |title=Near fixation of melanism in leopards of the Malay Peninsula |journal=Journal of Zoology |year=2010 |volume=282 |issue=3 |pages=201–206 |doi=10.1111/j.1469-7998.2010.00731.x}}{{cite journal |author1=Da Silva L. G. |author2=Kawanishi, K. |first=K. |author3=Henschel P. |author4=Kittle, A. |author5=Sanei, A. |author6=Reebin, A. |author7=Miquelle, D. |author8=Stein, A. B. |author9=Watson, A. |author10=Kekule, L. B. |author11=Machado, R. B. |author12=Eizirik, E. |name-list-style=amp |year=2017 |title=Mapping black panthers: Macroecological modeling of melanism in leopards (Panthera pardus) |journal= PLOS ONE |volume=12 |issue=4 |pages=e0170378 |doi=10.1371/journal.pone.0170378 |pmid=28379961 |pmc=5381760 |bibcode=2017PLoSO..1270378D |doi-access=free}}

In India, nine pale and white leopards were reported between 1905 and 1967.{{cite journal |author=Divyabhanusinh |year=1993 |title=On mutant leopards Panthera pardus from India |journal=Journal of the Bombay Natural History Society |volume=90 |issue=1 |pages=88−89 |url=https://archive.org/details/journalofbombay901993bomb/page/88}}

Leopards exhibiting erythrism were recorded between 1990 and 2015 in South Africa's Madikwe Game Reserve and in Mpumalanga. The cause of this morph known as a "strawberry leopard" or "pink panther" is not well understood.{{Cite journal |last1=Pirie |first1=T. J. |last2=Thomas |first2=R. L. |last3=Fellowes |first3=M. D. E. |name-list-style=amp |date=2016 |title=Erythristic leopards Panthera pardus in South Africa |journal=Bothalia |volume=46 |issue=1 |pages=1–5 |doi=10.4102/abc.v46i1.2034 |doi-access=free}}

The leopard is a slender and muscular cat, with relatively short limbs and a broad head. It is sexually dimorphic with males larger and heavier than females. Males stand {{cvt|60|-|70|cm}} at the shoulder, while females are {{cvt|57|-|64|cm}} tall. The head-and-body length ranges between {{cvt|92|and|183|cm}} with a {{cvt|66|to|102|cm|}} long tail. Sizes vary geographically. Males typically weigh {{cvt|30.9|-|72|kg}}, and females {{cvt|20.5|-|43|kg}}.{{cite book |author1=Hunter, L. |author2=Henschel, P. |author3=Ray, J. C. |name-list-style=amp |chapter=Panthera pardus Leopard |pages=159–168 |date=2013 |isbn=978-1-4081-8996-2 |chapter-url=https://books.google.com/books?id=B_07noCPc4kC&pg=RA4-PA159 |editor1=Kingdon, J. |editor2=Happold, D. |editor3=Butynski, T. |editor4=Hoffmann, M. |editor5=Happold, M. |editor6=Kalina, J. |title=Mammals of Africa |publisher=Bloomsbury Publishing |location=London |access-date=2021-05-19 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145933/https://books.google.com/books?id=B_07noCPc4kC&pg=RA4-PA159#v=onepage&q&f=false |url-status=live}} Occasionally, large males can grow up to {{cvt|91|kg}}. Leopards from the Cape Province in South Africa are generally smaller, reaching only {{cvt|20|-|45|kg}} in males.{{cite book |last1=Nowak |first1=R. M. |title=Walker's Mammals of the World |year=1999 |publisher=Johns Hopkins University Press |location=Baltimore, US |isbn=978-0-8018-5789-8 |edition=Sixth |pages=828–831 |chapter=Panthera pardus (Leopard) |chapter-url=https://books.google.com/books?id=T37sFCl43E8C&pg=PA828 |access-date=2021-05-19 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145930/https://books.google.com/books?id=T37sFCl43E8C&pg=PA828#v=onepage&q&f=false |url-status=live}}

The heaviest wild leopard in Southern Africa weighed around {{cvt|96|kg}}, and it measured {{cvt|262|cm}}.{{cite book |editor-last1=Burnie |editor-first1=D. |editor-last2=Wilson |editor-first2=D. E. |name-list-style=amp |year=2001 |title=Animal: The Definitive Visual Guide to the World's Wildlife |publisher=DK Adult |isbn=978-0-7894-7764-4}} In 2016, an Indian leopard killed in Himachal Pradesh measured {{cvt|261|cm}} with an estimated weight of {{cvt|78.5|kg}}; it was perhaps the largest known wild leopard in India.{{cite news |year=2016 |title=Is this the longest leopard in India? |newspaper=The Times of India |url=https://timesofindia.indiatimes.com/city/chandigarh/Is-this-the-longest-leopard-in-India/articleshow/56227308.cms |access-date=2018-03-14 |archive-date=2019-04-06 |archive-url=https://web.archive.org/web/20190406154410/https://timesofindia.indiatimes.com/city/chandigarh/Is-this-the-longest-leopard-in-India/articleshow/56227308.cms |url-status=live}}

The largest recorded skull of a leopard was found in India in 1920 and measured {{cvt|28|cm}} in basal length, {{cvt|20|cm}} in breadth, and weighed {{cvt|1|kg}}. The skull of an African leopard measured {{cvt|286|mm}} in basal length, and {{cvt|181|mm}} in breadth, and weighed {{cvt|790|g}}.{{cite journal |author=Prater, S. H. |title=Record panther skull (P. p. pardus) |journal=The Journal of the Bombay Natural History Society |date=1921 |volume=XXVII |issue=Part IV |pages=933–935 |url=https://www.biodiversitylibrary.org/item/95740#page/1055/mode/1up |access-date=2019-08-24 |archive-date=2019-05-02 |archive-url=https://web.archive.org/web/20190502064501/https://www.biodiversitylibrary.org/item/95740#page/1055/mode/1up |url-status=live}}

File:David Raju Leopard0827.jpg

File:Edgar Grooming (49611421486).jpg]]

The leopard has the largest distribution of all wild cats, occurring widely in Africa and Asia, although populations are fragmented and declining. It inhabits foremost savanna and rainforest, and areas where grasslands, woodlands and riparian forests remain largely undisturbed. It also persists in urban environments, if it is not persecuted, has sufficient prey and patches of vegetation for shelter during the day.{{Cite journal |last1=Powell |first1=J. |last2=Axmacher |first2=J. C. |last3=Linnell |first3=J. D.C. |last4=Durant |first4=S. M. |title=Diverse locations and a long history: Historical context for urban Leopards (Panthera pardus) in the early anthropocene From Seoul, Korea |year=2021 |doi=10.3389/fcosc.2021.765911 |journal=Frontiers in Conservation Science |volume=2 |page=765911 |doi-access=free |name-list-style=amp |hdl=11250/2999764 |hdl-access=free}}

The leopard's range in West Africa is estimated to have drastically declined by 95%, and in the Sahara desert by 97%.{{cite journal |last1=Jacobson|first1=A. P. |last2=Gerngross |first2=P. |last3=Lemeris |first3=J. R. Jr.|last4=Schoonover |first4=R. F. |name-list-style=amp |last5=Anco |first5=C. |last6=Breitenmoser-Würsten |first6=C. |last7=Durant |first7=S. M. |last8=Farhadinia |first8=M. S. |last9=Henschel |first9=P. |last10=Kamler |first10=J. F. |last11=Laguardia |first11=A. |last12=Rostro-García |first12=S. |last13=Stein |first13=A. B. |last14=Dollar |first14=L. |title=Leopard (Panthera pardus) status, distribution, and the research efforts across its range |journal=PeerJ |date=2016 |volume=4 |pages=e1974 |doi=10.7717/peerj.1974 |pmid=27168983 |pmc=4861552 |doi-access=free}} In sub-Saharan Africa, it is still numerous and surviving in marginal habitats where other large cats have disappeared.{{Cite journal |last1=Pirie |first1=T. J. |last2=Thomas |first2=R. L. |last3=Fellowes |first3=M. D. E. |name-list-style=amp |date=2017 |title=Increasing game prices may alter farmers' behaviours towards leopards (Panthera pardus) and other carnivores in South Africa |page=e3369 |journal=PeerJ |volume=5 |doi=10.7717/peerj.3369|pmid=28584709 |pmc=5452990 |doi-access=free}} In southeastern Egypt, an individual found killed in 2017 was the first sighting of the leopard in this area in 65 years.{{Cite journal |last1=Soultan, A. |last2=Attum, O. |last3=Hamada, A. |last4=Hatab, E. B. |last5=Ahmed, S. E. |last6=Eisa, A. |last7=Al Sharif, I. |last8=Nagy, A. |name-list-style=amp |author9=Shohdi, W. |date=2017 |title=Recent observation for leopard Panthera pardus in Egypt |journal=Mammalia |volume=81 |issue=1 |pages=115–117 |doi=10.1515/mammalia-2015-0089 |s2cid=90676105}}

In West Asia, the leopard inhabits remain in the areas of southern and southeastern Anatolia.{{cite journal |author1=Toyran, K. |title=Noteworthy record of Panthera pardus in Turkey (Carnivora: Felidae) |journal=Fresenius Environmental Bulletin |year=2018 |volume=27 |issue=11 |pages=7348–7353}}{{cite journal |author1=Karataş, A. |name-list-style=amp |author2=Bulut, Ş. |author3=Akbaba, B. |year=2021 |title=Camera trap records confirm the survival of the Leopard (Panthera pardus L., 1758) in eastern Turkey (Mammalia: Felidae) |journal=Zoology in the Middle East |volume= 67 |issue= 3|pages=198–205 |s2cid=235564429 |doi=10.1080/09397140.2021.1924419}}{{Cite journal |title=New records of the living Anatolian Leopard (Panthera pardus tulliana L., 1758) in the Mediterranean region of Turkey |author1=Ünal, Y. |author2=Uysal, H. |author3=Koca, A. |author4=Zenbilci, M. |journal=Applied Ecology and Environmental Research |volume=21 |pages=1043–1059 |issue=2 |doi=10.15666/aeer/2102_10431059 |date=2023 |doi-access=free |name-list-style=amp}}

Leopard populations in the Arabian Peninsula are small and fragmented.{{cite journal |author1=Spalton, J. A. |author2=Al Hikmani, H. M. |name-list-style=amp |year=2006 |title=The Leopard in the Arabian Peninsula – Distribution and subspecies status |journal=Cat News |issue=Special Issue 1 |pages=4–8 |url=http://catsg.org/fileadmin/filesharing/5.Cat_News/5.3._Special_Issues/5.3.1._SI_1/Spalton___Al_Hikmani_2006_Leopard_in_the_Arabian_Peninsula.pdf }}{{cite journal |last1=Judas |first1=J. |last2=Paillat |first2=P. |last3=Khoja |first3=A. |last4=Boug |first4=A. |name-list-style=amp |year=2006 |title=Status of the Arabian leopard in Saudi Arabia |journal=Cat News |issue=Special Issue 1 |pages=11–19 |url=http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.2._Status_Reports/leopard/Judas_et_al_2006_Status_of_the_Arabian_Leopard_in_Saudi_Arabia.pdf }}{{cite journal |last1=Al Jumaily, M. |last2=Mallon, D. P. |last3=Nasher, A. K. |last4=Thowabeh, N. |name-list-style=amp |year=2006 |title=Status Report on Arabian Leopard in Yemen |journal=Cat News |issue=Special Issue 1 |pages=20–25 |url=http://catsg.org/fileadmin/filesharing/5.Cat_News/5.3._Special_Issues/5.3.1._SI_1/Jumaily_et_al_2006_Status_Report_on_Arabian_Leopard_in_Yemen.pdf}}

In the Indian subcontinent, the leopard is still relatively abundant, with greater numbers than those of other Panthera species. Some leopard populations in India live quite close to human settlements and even in semi-developed areas. Although adaptable to human disturbances, leopards require healthy prey populations and appropriate vegetative cover for hunting for prolonged survival and thus rarely linger in heavily developed areas. Due to the leopard's stealth, people often remain unaware that it lives in nearby areas.{{cite web |author=Arthreya, V. |year=2012 |title=Living with Leopards Outside Protected Areas in India |url=http://www.conservationindia.org/articles/living-with-leopards-outside-protected-areas-in-india |publisher=Conservation India |access-date=2012-12-10 |archive-date=2012-12-21 |archive-url=https://web.archive.org/web/20121221051725/http://www.conservationindia.org/articles/living-with-leopards-outside-protected-areas-in-india |url-status=live}} As of 2020, the leopard population within forested habitats in India's tiger range landscapes was estimated at 12,172 to 13,535 individuals. Surveyed landscapes included elevations below {{cvt|2600|m}} in the Shivalik Hills and Gangetic plains, Central India and Eastern Ghats, Western Ghats, the Brahmaputra River basin and hills in Northeast India.{{cite report |author=Jhala, Y.V. |author-link=Yadvendradev Vikramsinh Jhala |author2=Qureshi, Q. |author3=Yadav, S.P. |name-list-style=amp |year=2020 |title=Status of leopards in India, 2018. Technical Report TR/2020/16 |location=New Delhi and Dehradun |publisher=National Tiger Conservation Authority, Government of India and Wildlife Institute of India}}

In Nepal's Kanchenjunga Conservation Area, a melanistic leopard was photographed at an elevation of {{cvt|4300|m}} by a camera trap in May 2012.{{cite journal |author1=Thapa, K. |author2=Pradhan, N. M. B. |author3=Berker, J. |author4=Dhakal, M. |author5=Bhandari, A. R. |author6=Gurung, G. S. |author7=Rai, D. P. |author8=Thapa, G. J. |author9=Shrestha, S. |author10=Singh, G. R. |name-list-style=amp |year=2013 |title=High elevation record of a leopard cat in the Kangchenjunga Conservation Area, Nepal |journal=Cat News |issue=58 |pages=26–27}}

In Sri Lanka, leopards were recorded in Yala National Park and in unprotected forest patches, tea estates, grasslands, home gardens, pine and eucalyptus plantations.{{cite journal |author=Kittle, A. M. |author2=Watson, A. C. |author3=Chanaka Kumara, P. H. |author4=Nimalka Sanjeewani, H. K. |name-list-style=amp |year=2014 |title=Status and distribution of the leopard in the central hills of Sri Lanka |journal=Cat News |issue=56 |pages=28−31}}{{cite journal |last1=Kittle |first1=A. M. |last2=Watson |first2=A. C. |last3=Kumara |first3=P. H. S. C. |last4=Sandanayake |first4=S. D. K. C. |last5=Sanjeewani |first5=H. K. N. |last6=Fernando |first6=T. S. P. |name-list-style=amp |year=2014 |title=Notes on the diet and habitat selection of the Sri Lankan Leopard Panthera pardus kotiya (Mammalia: Felidae) in the central highlands of Sri Lanka |journal=Journal of Threatened Taxa |volume=6 |issue=9 |pages=6214–6221 |doi=10.11609/JoTT.o3731.6214-21 |doi-access=free}}

In Myanmar, leopards were recorded for the first time by camera traps in the hill forests of Myanmar's Karen State.{{cite journal |author1=Saw Sha Bwe Moo |author2=Froese, G.Z.L. |author3=Gray, T.N.E. |name-list-style=amp |year=2017 |title=First structured camera-trap surveys in Karen State, Myanmar, reveal high diversity of globally threatened mammals |journal=Oryx |volume=52 |issue=3 |pages=537−543 |doi=10.1017/S0030605316001113 |doi-access=free}} The Northern Tenasserim Forest Complex in southern Myanmar is considered a leopard stronghold. In Thailand, leopards are present in the Western Forest Complex, Kaeng Krachan-Kui Buri, Khlong Saeng-Khao Sok protected area complexes and in Hala Bala Wildlife Sanctuary bordering Malaysia. In Peninsular Malaysia, leopards are present in Belum-Temengor, Taman Negara and Endau-Rompin National Parks.{{cite journal | last1=Rostro-García |first1=S. |last2=Kamler |first2 =J. F. |last3=Ash |first3=E. |last4=Clements |first4=G. R. |last5=Gibson |first5=L. |last6=Lynam |first6=A. J. |last7=McEwin |first7=R. |last8=Naing |first8=H. |last9=Paglia |first9=S. |name-list-style=amp |year=2016 |title=Endangered leopards: Range collapse of the Indochinese leopard (Panthera pardus delacouri) in Southeast Asia |journal=Biological Conservation |volume=201 |pages=293–300 |doi=10.1016/j.biocon.2016.07.001|bibcode=2016BCons.201..293R |hdl=10722/232870 |hdl-access=free}}

In Laos, leopards were recorded in Nam Et-Phou Louey National Biodiversity Conservation Area and Nam Kan National Protected Area.{{cite journal |author=Johnson, A. |author2=Vongkhamheng, C. |name-list-style=amp |author3=Hedemark, M. |author4=Saithongdam, T. |year=2006 |title=Effects of human–carnivore conflict on tiger (Panthera tigris) and prey populations in Lao PDR |journal=Animal Conservation |volume=9 |issue=4 |pages=421–430 |doi=10.1111/j.1469-1795.2006.00049.x |bibcode=2006AnCon...9..421J |s2cid=73637721 |url=http://faculty.nelson.wisc.edu/treves/pubs/Arlyne/animal_cons.pdf |archive-url=https://web.archive.org/web/20170810101758/http://faculty.nelson.wisc.edu/treves/pubs/Arlyne/animal_cons.pdf |archive-date=2017-08-10 |url-status=live}}{{cite book |author=Robichaud, W. |name-list-style=amp |author2=Insua-Cao |author3=Sisomphane, P. C. |author4=Chounnavanh, S. |year=2010 |title=A scoping mission to Nam Kan National Protected Area, Lao PDR |publisher=Fauna & Flora International |chapter=Appendix 4 |pages=33−42 |chapter-url=https://studylib.net/doc/18697244/a-scoping-mission-to-nam-kan-national-protected-area--lao... |access-date=2019-04-07 |archive-date=2019-04-07 |archive-url=https://web.archive.org/web/20190407154414/https://studylib.net/doc/18697244/a-scoping-mission-to-nam-kan-national-protected-area--lao... |url-status=live}}

In Cambodia, leopards inhabit deciduous dipterocarp forest in Phnom Prich Wildlife Sanctuary and Mondulkiri Protected Forest.{{cite journal |last1=Gray |first1=T. N. |last2=Phan |first2=C. |name-list-style=amp |year=2011 |title=Habitat preferences and activity patterns of the larger mammal community in Phnom Prich Wildlife Sanctuary, Cambodia |journal=The Raffles Bulletin of Zoology |volume=59 |issue=2 |pages=311−318 |url=https://www.researchgate.net/publication/228843849 |access-date=2019-04-16 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505150044/https://www.researchgate.net/publication/228843849_Habitat_preferences_and_activity_patterns_of_the_larger_mammal_community_in_Phnom_Prich_Wildlife_Sanctuary_Cambodia |url-status=live}}{{cite journal |last1=Gray |first1=T. N. E. |year=2013 |title=Activity patterns and home ranges of Indochinese leopard Panthera pardus delacouri in the Eastern Plains Landscape, Cambodia |journal=Natural History Bulletin of the Siam Society |volume=59 |pages=39−47 |url=http://www.siamese-heritage.org/nhbsspdf/vol051-060/NHBSS_059_1h_Gray_ActivityPatternsAndH.pdf |archive-url=https://web.archive.org/web/20160222060912/http://www.siamese-heritage.org/nhbsspdf/vol051-060/NHBSS_059_1h_Gray_ActivityPatternsAndH.pdf |archive-date=2016-02-22 |url-status=live}}

In southern China, leopards were recorded only in the Qinling Mountains during surveys in 11 nature reserves between 2002 and 2009.{{cite journal |author=Li, S. |author2=Wang, D. |author3=Lu, Z. |author4=Mc Shea, W. J. |name-list-style=amp |year=2010 |title=Cats living with pandas: The status of wild felids within giant panda range, China |journal=Cat News |volume=52 |pages=20–23}}

In Java, leopards inhabit dense tropical rainforests and dry deciduous forests at elevations from sea level to {{cvt|2540|m}}. Outside protected areas, leopards were recorded in mixed agricultural land, secondary forest and production forest between 2008 and 2014.{{cite journal |last1=Wibisono |first1=H. T. |last2=Wahyudi |first2=H. A. |last3=Wilianto |first3=E. |last4=Pinondang |first4=I. M. R. |last5=Primajati |first5=M. |last6=Liswanto |first6=D. |last7=Linkie |first7=M. |name-list-style=amp |year=2018 |title=Identifying priority conservation landscapes and actions for the Critically Endangered Javan leopard in Indonesia: Conserving the last large carnivore in Java Island |journal=PLOS ONE |volume=13 |issue=6 |page=e0198369 |doi=10.1371/journal.pone.0198369 |pmid=29949588 |pmc=6021038 |bibcode=2018PLoSO..1398369W |doi-access=free}}

In the Russian Far East, it inhabits temperate coniferous forests where winter temperatures reach a low of {{cvt|−25|°C|°F}}.

The leopard is a solitary and territorial animal. It is typically shy and alert when crossing roadways and encountering oncoming vehicles, but may be emboldened to attack people or other animals when threatened. Adults associate only in the mating season. Females continue to interact with their offspring even after weaning and have been observed sharing kills with their offspring when they can not obtain any prey. They produce a number of vocalizations, including growls, snarls, meows, and purrs. Cubs call their mother with an urr-urr sound. The most notable vocalization is the 'sawing' roar, which consists of deep, repeated strokes. This likely functions in establishing territories and attracting mates.{{cite journal|last1=Growcott|first1=J|last2=Lobora|first2=A|last3=Markham|first3=A|last4=Searle|first4=C. E.|last5=Wahlström|first5=J|last6=Wijers|first6=M|last7=Simmons|first7=B. I.|year=2024| title=The secret acoustic world of leopards: A paired camera trap and bioacoustics survey facilitates the individual identification of leopards via their roars|journal=Remote Sensing in Ecology and Conservation|doi=10.1002/rse2.429}}

The whitish spots on the back of its ears are thought to play a role in communication.{{cite book |last1=Leyhausen |first1=P. |year=1979 |title=Cat behavior: the predatory and social behavior of domestic and wild cats |location=Berlin |publisher=Garland Publishing, Incorporated |page=281 |isbn=9780824070175}}

It has been hypothesized that the white tips of their tails may function as a 'follow-me' signal in intraspecific communication. However, no significant association were found between a conspicuous colour of tail patches and behavioural variables in carnivores.{{cite journal |last1=Ortolani |first1=A. |year=1999 |title=Spots, stripes, tail tips and dark eyes: predicting the function of carnivore colour patterns using the comparative method |journal=Biological Journal of the Linnean Society |volume=67 |issue=4 |pages=433–476 |doi=10.1111/j.1095-8312.1999.tb01942.x |doi-access=free}}{{cite journal |last1=Caro |first1=T. |year=2005 |title=The adaptive significance of coloration in mammals |journal=BioScience |volume=55 |issue=2 |pages=125–136 |doi=10.1641/0006-3568(2005)055[0125:TASOCI]2.0.CO;2 |doi-access=free}}

Leopards are mainly active from dusk till dawn and will rest for most of the day and some hours at night in thickets, among rocks or over tree branches. Leopards have been observed walking {{cvt|1|-|25|km|mi}} across their range at night; wandering up to {{cvt|75|km}} if disturbed. In some regions, they are nocturnal.{{cite journal |last1=Hunter |first1=L. |last2=Balme |first2=G. |last3=Walker |first3=C. |last4=Pretorius |first4=K. |last5=Rosenberg |first5=K. |name-list-style=amp |year=2003 |title=The landscape ecology of leopards (Panthera pardus) in northern KwaZulu-Natal, South Africa: a preliminary project report |journal=Ecological Journal |volume=5 |pages=24–30 |url=http://www.biolsci.monash.edu.au/research/leopards/docs/ecojournal-2003.pdf |url-status=dead |archive-url=https://web.archive.org/web/20090304233241/http://www.biolsci.monash.edu.au/research/leopards/docs/ecojournal-2003.pdf |archive-date=March 4, 2009}} {{open access}}{{cite journal |last1=Spalton |first1=J.A. |name-list-style=amp |last2=Al Hikmani |first2=H. M. |last3=Willis |first3=D. |last4=Said |first4=A. S. B. |title=Critically endangered Arabian leopards Panthera pardus nimr persist in the Jabal Samhan Nature Reserve, Oman |journal=Oryx |date=2006 |volume=40 |issue=3 |pages=287–294 |doi=10.1017/S0030605306000743 |doi-access=free}} In western African forests, they have been observed to be largely diurnal and hunting during twilight, when their prey animals are active; activity patterns vary between seasons.{{cite journal |last1=Jenny |first1=D. |last2=Zuberbuhler |first2=K. |name-list-style=amp |title=Hunting behaviour in west African forest leopards |journal=African Journal of Ecology |date=2005 |volume=43 |issue=3 |pages=197–200 |doi=10.1111/j.1365-2028.2005.00565.x |bibcode=2005AfJEc..43..197J |url=https://www.researchgate.net/publication/229882994 |access-date=2018-09-25 |archive-date=2018-09-25 |archive-url=https://web.archive.org/web/20180925142137/https://www.researchgate.net/publication/229882994 |url-status=live}}

Leopards can climb trees quite skillfully, often resting on tree branches and descending headfirst.

They can run at over {{cvt|58|km/h|mph m/s}}, leap over {{cvt|6|m}} horizontally, and jump up to {{cvt|3|m}} vertically.{{Cite book |last1=Sunquist |first1=M. E. |year=2002 |last2=Sunquist |first2=F. |name-list-style=amp |title=Wild Cats of the World |publisher=University of Chicago Press |location=Chicago |isbn=978-0-226-77999-7 |chapter=Leopard Panthera pardus |pages=318–342 |chapter-url=https://books.google.com/books?id=IF8nDwAAQBAJ&pg=PA320 |access-date=2021-05-19 |archive-date=2024-04-30 |archive-url=https://web.archive.org/web/20240430033051/https://books.google.com/books?id=IF8nDwAAQBAJ&pg=PA320#v=onepage&q&f=false |url-status=live}}

{{image frame|align=center|border=no|content=

leopard rear view soft.jpg|A female leopard with white markings on the backs of her ears

leopard walking.jpg|A female leopard showing white spots on the tail

Leopard davidraju 68.jpg|A leopard climbing down a tree

Leopard hunting a bush pig - DPLA - 57da78c992bc6073d2751f3f8936aad0.jpg|A leopard hunting a bushpig

Leopard.ogv|Video of a leopard in the wild

}}

File:Day 47 Leopard (Panthera pardus) male marking a bush with urine ... (53310719953).jpg his territory{{Cite book |last1=Bothma |first1=Jacobus du P. |url=https://books.google.com/books?id=DPvuCAAAQBAJ |title=Larger Carnivores of the African Savannas |last2=Walker |first2=Clive |date=2013-11-11 |publisher=Springer Science & Business Media |isbn=978-3-662-03766-9 |language=en |access-date=2024-01-06 |archive-date=2024-01-06 |archive-url=https://web.archive.org/web/20240106170223/https://books.google.com/books?id=DPvuCAAAQBAJ |url-status=live}}]]

In Kruger National Park, most leopards tend to keep {{cvt|1|km|mi}} apart.{{cite book |last=Bailey |first=T. N. |year=1993 |title=The African leopard: a study of the ecology and behaviour of a solitary felid |publisher=Columbia University Press |location=New York |isbn=978-1-932846-11-9}} Males occasionally interact with their partners and cubs, and exceptionally this can extend beyond to two generations.{{Cite journal |last1=Pirie |first1=T. J.|last2=Thomas |first2=R. L.|last3=Reilly|first3=B. K. |last4=Fellowes |first4=M. D. E. |name-list-style=amp |date=2014 |title=Social interactions between a male leopard (Panthera pardus) and two generations of his offspring |journal=African Journal of Ecology |volume=52 |issue=4 |pages=574–576 |doi=10.1111/aje.12154 |doi-access=free |bibcode=2014AfJEc..52..574P}} Aggressive encounters are rare, typically limited to defending territories from intruders. In a South African reserve, a male was wounded in a male–male territorial battle over a carcass.

Males occupy home ranges that often overlap with a few smaller female home ranges, probably as a strategy to enhance access to females. In the Ivory Coast, the home range of a female was completely enclosed within a male's.{{cite journal |author=Jenny, D. |year=1996 |title=Spatial organization of leopards Panthera pardus in Tai National Park, Ivory Coast: Is rainforest habitat a "tropical haven"? |journal=Journal of Zoology |issue=3 |volume=240 |pages=427–440 |doi=10.1111/j.1469-7998.1996.tb05296.x |url=https://www.researchgate.net/publication/227901726 |access-date=2021-09-06 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145917/https://www.researchgate.net/publication/227901726_Spatial_organization_of_leopards_Panthera_pardus_in_Tai_National_Park_Ivory_Coast_Is_rainforest_habitat_a_%27tropical_haven%27 |url-status=live}} Females live with their cubs in home ranges that overlap extensively, probably due to the association between mothers and their offspring. There may be a few other fluctuating home ranges belonging to young individuals. It is not clear if male home ranges overlap as much as those of females do. Individuals try to drive away intruders of the same sex.

A study of leopards in the Namibian farmlands showed that the size of home ranges was not significantly affected by sex, rainfall patterns or season; the higher the prey availability in an area, the greater the leopard population density and the smaller the size of home ranges, but they tend to expand if there is human interference.{{cite journal|last1=Marker |first1=L. L. |last2=Dickman |first2=A. J. |name-list-style=amp |title=Factors affecting leopard (Panthera pardus) spatial ecology, with particular reference to Namibian farmlands |journal=South African Journal of Wildlife Research |date=2005 |volume=35 |issue=2 |pages=105–115 |url=http://reference.sabinet.co.za/webx/access/journal_archive/03794369/3418.pdf}} {{open access}}

Sizes of home ranges vary geographically and depending on habitat and availability of prey. In the Serengeti, males have home ranges of {{cvt|33|-|38|km2|sqmi}} and females of {{cvt|14|-|16|km2|sqmi}};{{cite journal |last1=Bertram |first1=B. C. R.|title=Leopard ecology as studied by radio tracking |journal=Symposia of the Zoological Society of London |date=1982 |volume=49|pages=341–352}}{{cite journal |last1=Mizutani, F. |last2=Jewell, P. A. |name-list-style=amp |year=1998 |title=Home-range and movements of leopards (Panthera pardus) on a livestock ranch in Kenya |journal=Journal of Zoology |pages=269–286 |volume=244 |issue=2 |doi=10.1017/S0952836998002118}} but males in northeastern Namibia of {{cvt|451|km2|sqmi}} and females of {{cvt|188|km2|sqmi}}.{{cite journal |last1=Stander |first1=P. E. |last2=Haden |first2=P. J. |last3=Kaqece |first3=II. |last4=Ghau |first4=II. |name-list-style=amp|title=The ecology of asociality in Namibian leopards|journal=Journal of Zoology |date=1997 |volume=242|issue=2 |pages=343–364 |doi=10.1111/j.1469-7998.1997.tb05806.x}} They are even larger in arid and montane areas. In Nepal's Bardia National Park, male home ranges of {{cvt|48|km2|sqmi}} and female ones of {{cvt|5|-|7|km2|sqmi}} are smaller than those generally observed in Africa.{{cite journal |author=Odden, M. |author2=Wegge, P. |name-list-style=amp |year=2005 |title=Spacing and activity patterns of leopards Panthera pardus in the Royal Bardia National Park, Nepal |journal=Wildlife Biology |volume=11 |issue=2 |pages=145–152 |doi=10.2981/0909-6396(2005)11[145:SAAPOL]2.0.CO;2 |doi-access=free |s2cid=86140708}}

{{multiple image |perrow=1 |image1=Leopard (Panthera pardus pardus) stalking.jpg |caption1=Leopard stalking |image2=Leopard kill - KNP - 001.jpg|caption2=Leopard applying a throat bite to a bushbuck|image3=Leopardo (Panthera pardus) devorando un antílope, parque nacional Kruger, Sudáfrica, 2018-07-26, DD 06.jpg |caption3=Leopard caches a kill in a tree}}

The leopard is a carnivore that prefers medium-sized prey with a body mass ranging from {{cvt|10|–|40|kg|lb}}. Prey species in this weight range tend to occur in dense habitat and to form small herds. Species that prefer open areas and have well-developed anti-predator strategies are less preferred. More than 100 prey species have been recorded. The most preferred species are ungulates, such as impala, bushbuck, common duiker and chital. Primates preyed upon include white-eyelid mangabeys, guenons and gray langurs. Leopards also kill smaller carnivores like black-backed jackal, bat-eared fox, genet and cheetah.{{cite journal |last1=Hayward |first1=M.W. |last2=Henschel |first2=P. |last3=O'Brien |first3=J. |last4=Hofmeyr |first4=M. |last5=Balme |first5=G. |last6=Kerley |first6=G. I. H. |name-list-style=amp |title=Prey preferences of the leopard (Panthera pardus) |journal=Journal of Zoology |date=2006 |volume=270 |issue=4 |pages=298–313 |doi=10.1111/j.1469-7998.2006.00139.x |url=http://www.ibs.bialowieza.pl/g2/pdf/1596.pdf |archive-url=https://web.archive.org/web/20121105063845/http://www.ibs.bialowieza.pl/g2/pdf/1596.pdf |archive-date=2012-11-05 |url-status=live}} In urban environments, domestic dogs provide an important food source. The largest prey killed by a leopard was reportedly a male eland weighing {{cvt|900|kg}}. A study in Wolong National Nature Reserve in southern China demonstrated variation in the leopard's diet over time; over the course of seven years, the vegetative cover receded, and leopards opportunistically shifted from primarily consuming tufted deer to pursuing bamboo rats and other smaller prey.{{cite journal |author1=Johnson, K. G. |author2=Wei, W. |author3=Reid, D. G. |author4=Jinchu, H. |date=1993 |title=Food habits of Asiatic leopards (Panthera pardus fusca) in Wolong Reserve, Sichuan, China |journal=Journal of Mammalogy |volume=74 |issue=3 |pages=646–650 |jstor=1382285 |doi=10.2307/1382285}}

The leopard depends mainly on its acute senses of hearing and vision for hunting.{{cite book |last1=Mills |first1=M. G. L. |last2=Hes |first2=L. |name-list-style=amp |isbn=978-0-947430-55-9|pages=178–180 |publisher=Struik Publishers|location=Cape Town, South Africa |title=The Complete Book of Southern African Mammals |year=1997}} It primarily hunts at night in most areas. In western African forests and Tsavo National Park, they have also been observed hunting by day.{{cite thesis |last=Hamilton |first=P. H. |year=1976 |title=The movements of leopards in Tsavo National Park, Kenya, as determined by radio-tracking |degree= M.Sc. |publisher=University of Nairobi |location=Nairobi}} They usually hunt on the ground. In the Serengeti, they have been seen to ambush prey by descending on it from trees.{{cite journal |last1=Kruuk |first1=H. |last2=Turner |first2=M. |name-list-style=amp |date=1967 |title=Comparative notes on predation by lion, leopard, cheetah and wild dog in the Serengeti area, East Africa |journal=Mammalia |volume=31 |issue=1 |pages=1–27 |doi=10.1515/mamm.1967.31.1.1 |s2cid=84619500}} It stalks its prey and tries to approach as closely as possible, typically within {{cvt|5|m}} of the target, and, finally, pounces on it and kills it by suffocation. It kills small prey with a bite to the back of the neck, but holds larger animals by the throat and strangles them. It caches kills up to {{cvt|2|km|mi}} apart. It is able to take large prey due to its powerful jaw muscles, and is therefore strong enough to drag carcasses heavier than itself up into trees; an individual was seen to haul a young giraffe weighing nearly {{cvt|125|kg}} up {{cvt|5.7|m|ftin}} into a tree. It eats small prey immediately, but drags larger carcasses over several hundred metres and caches it safely in trees, bushes or even caves; this behaviour allows the leopard to store its prey away from rivals, and offers it an advantage over them. The way it stores the kill depends on local topography and individual preferences, varying from trees in Kruger National Park to bushes in the plain terrain of the Kalahari.{{cite book |last=Schaller |first=G. |year=1972 |title=Serengeti: a kingdom of predators |publisher=Knopf |location=New York |isbn=978-0-394-47242-3 |url=https://books.google.com/books?id=wJ_wAAAAMAAJ |access-date=2020-09-18 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505150524/https://books.google.com/books?id=wJ_wAAAAMAAJ |url-status=live}}

Average daily consumption rates of {{cvt|3.5|kg|lboz}} were estimated for males and of {{cvt|2.8|kg|lboz}} for females. In the southern Kalahari Desert, leopards meet their water requirements by the bodily fluids of prey and succulent plants; they drink water every two to three days and feed infrequently on moisture-rich plants such as gemsbok cucumbers, watermelon and Kalahari sour grass.{{cite journal| last1=Bothma |first1=J. du P. |title=Water-use by southern Kalahari leopards |journal=South African Journal of Wildlife Research |date=2005 |volume=35 |pages=131–137 |url=http://reference.sabinet.co.za/webx/access/journal_archive/03794369/2353.pdf}} {{open access}}

File:Lioness vs Leopard 9 July 2016 Latest Sightings 1.png

Across its range, the leopard coexists with a number of other large predators. In Africa, it is part of a large predator guild with lions, cheetahs, spotted and brown hyenas, and African wild dogs.{{cite journal|last1=Rafiq|first1=K.|last2=Wayward |first2=M. W.|last3=Wilson|first3=A. M.|last4=Meloro|first4=C. |last5=Jordan|first5=N. R. |last6=Wich |first6=S. A. |last7=McNutt|first7=J. W. |last8=Golabek|first8=K. A.|year=2020|title=Spatial and temporal overlaps between leopards (Panthera pardus) and their competitors in the African large predator guild|journal=Journal of Zoology |volume=311 |issue=4 |pages=246–259 |doi=10.1111/jzo.12781}} The leopard is dominant only over the cheetah while the others have the advantage of size, pack numbers or both. Lions pose a great mortal threat and can be responsible for 22% of leopard deaths in Sabi Sand Game Reserve. Spotted hyenas are less threatening but are more likely to steal kills, being the culprits of up to 50% of stolen leopard kills in the same area.{{cite journal|last1=Balme|first1=G. A.|last2=Pitman|first2=R. T.|last3=Robinson|first3=H. S.|last4=Miller|first4=J. R. B.|last5=Funston|first5=P. J.|last6=Hunter|first6=L. T. B.|year=2017|title=Leopard distribution and abundance is unaffected by interference competition with lions|journal=Behavioral Ecology |volume=28 |issue=5 |pages=1348–1358 |doi=10.1093/beheco/arx098}}{{cite journal|last1=Balme|first1=G. A. |last2=Miller |first2=J. R. B.|last3=Pitman|first3=R. T.|last4=Hunter|first4=L. T. B.|year=2017|title=Caching reduces kleptoparasitism in a solitary, large felid|journal=Journal of Animal Ecology|volume=86|issue=3 |pages=634–644 |doi=10.1111/1365-2656.12654 |pmid=28217865 |bibcode=2017JAnEc..86..634B}} To counter this, leopards store their kills in the trees and out of reach.{{cite journal|last1=Vissia|first1=S.|last2=Fattebert|first2=J. |last3=van Langevelde|first3=F. |year=2022 |title=Leopard density and interspecific spatiotemporal interactions in a hyena-dominated landscape|journal=Ecology and Evolution |volume=12 |issue=10 |page=e9365 |doi=10.1002/ece3.9365 |pmid=36225822 |pmc=9534747 |bibcode=2022EcoEv..12E9365V}} Lions have a high success rate in fetching leopard kills from trees. Leopards do not seem to actively avoid their competitors but rather difference in prey and habitat preferences appear to limit their spatial overlap. In particular, leopards use heavy vegetation regardless of whether lions are present in an area and both cats are active at the same time of day.{{cite journal |last1=Miller|first1=J. R. B. |last2=Pitman |first2=R. T. |last3=Mann |first3=G. K. H. |last4=Fuller|first4=A. K.|last5=Balme|first5=G. A. |year=2018|title=Lions and leopards coexist without spatial, temporal or demographic effects of interspecific competition |journal=Journal of Animal Ecology |volume=87 |issue=6 |pages=1709–1726 |doi=10.1111/1365-2656.12883 |pmid=30010193 |bibcode=2018JAnEc..87.1709M}}

In Asia, the leopard's main competitors are tigers and dholes. Both the larger tiger and pack-living dhole dominate leopards during encounters. Interactions between the three predators involve chasing, stealing kills and direct killing.{{cite journal |last1=Srivathsa |first1=A. |last2=Ramachandran|first2=V. |last3=Saravanan |first3=P. |last4=Sureshbabu |first4=A. |last5=Ganguly |first5=D. |last6=Ramakrishnan |first6=U. |year=2023 |title=Topcats and underdogs: intraguild interactions among three apex carnivores across Asia's forestscapes |journal=Biological Reviews |volume=98 |issue=6 |pages=2114–2135 |doi=10.1111/brv.12998 |pmid=37449566 |s2cid=259903849 |name-list-style=amp}} Tigers appear to inhabit the deep parts of the forest while leopards and dholes are pushed closer to the fringes.{{cite journal |author=Thinley, P. |author2=Rajaratnam, R. |author3=Lassoie, J. P. |author4=Morreale, S. J. |author5=Curtis, P. D. |author6=Vernes, K. |author7=Leki Leki |author8=Phuntsho, S. |author9=Dorji, T. |author10=Dorji, P. |name-list-style=amp |year=2018 |title=The ecological benefit of tigers (Panthera tigris) to farmers in reducing crop and livestock losses in the eastern Himalayas: Implications for conservation of large apex predators |journal=Biological Conservation |volume=219 |pages=119–125 |doi=10.1016/j.biocon.2018.08.007 |doi-access=free}} The three predators coexist by hunting different sized prey.{{cite journal |last=Karanth |first=K. U. |author2=Sunquist, M. E. |name-list-style=amp |year=2000 |title=Behavioural correlates of predation by tiger (Panthera tigris), leopard (Panthera pardus) and dhole (Cuon alpinus) in Nagarahole, India | journal=Journal of Zoology |volume=250 |pages=255–265 |doi=10.1111/j.1469-7998.2000.tb01076.x |issue=2}} In Nagarhole National Park, the average size for a leopard kill was {{cvt|37.6|kg}} compared to {{cvt|91.5|kg}} for tigers and {{cvt|43.4|kg}} for dholes.{{cite journal |author1=Karanth, K. U. |author2=Sunquist, M. E. |name-list-style=amp |title=Prey selection by tiger, leopard and dhole in tropical forests |jstor=5647 |doi=10.2307/5647 |journal=Journal of Animal Ecology |volume=64 |issue=4 |year=1995 |pages=439–450|bibcode=1995JAnEc..64..439K}} At Kui Buri National Park, following a reduction in prey numbers, tigers continued to feed on favoured prey while leopards and dholes had to increase their consumption of small prey.{{cite journal |author1=Steinmetz, R. |name-list-style=amp |author2=Seuaturien, N. |author3=Intanajitjuy, P. |author4=Inrueang, P. |author5=Prempree, K. |year=2021 |title=The effects of prey depletion on dietary niches of sympatric apex predators in Southeast Asia |journal=Integrative Zoology |volume=16 |issue=1 |pages=19–32 |doi=10.1111/1749-4877.12461|pmid=32627329 }} Leopards can live successfully in tiger habitat when there is abundant food and vegetation cover.{{cite journal |author1=Odden, M. |name-list-style=amp |author2=Wegge, P. |author3=Fredriksen, T. |year=2010 |title=Do tigers displace leopards? If so, why? |journal=Ecological Research |volume=25 |issue=4 |pages=875–881 |doi=10.1007/s11284-010-0723-1 |bibcode=2010EcoR...25..875O}} Otherwise, they appear to be less common where tigers are numerous. The recovery of the tiger population in Rajaji National Park during the 2000s led to a reduction in leopard population densities.{{cite journal |last1=Harihar |first1=A. |last2=Pandav |first2=B. |last3=Goyal |first3=S. P. |year=2011 |title=Responses of leopard Panthera pardus to the recovery of a tiger Panthera tigris population |journal=Journal of Applied Ecology |volume=48 |issue=3 |pages=806–814 |doi=10.1111/j.1365-2664.2011.01981.x|doi-access=free |bibcode=2011JApEc..48..806H |name-list-style=amp}}

{{multiple image |perrow=1 |image2=Leopards mating.jpg |caption2=Leopards mating |image3=David Raju Leopard 3457 (cropped).jpg |caption3=Leopard cubs in tree}}

In some areas, leopards mate all year round. In Manchuria and Siberia, they mate during January and February. On average, females begin to breed between the ages of 2½ and three, and males between the ages of two and three. The female's estrous cycle lasts about 46 days, and she is usually in heat for 6–7 days.{{cite journal |doi=10.1111/j.1748-1090.1966.tb01746.x |author=Sadleir, R. |year=1966 |title=Notes on the Reproduction of the larger Felidae |journal=International Zoo Yearbook|volume=6 |pages=184–187}} Gestation lasts for 90 to 105 days.{{cite book |author=Hemmer, H. |year=1976 |chapter=Gestation period and postnatal development in felids |pages=143–165 |editor=Eaton, R.L. |title=The World's Cats |volume=3 |publisher=Carnivore Research Institute, Univ. Washington, Seattle}} Cubs are usually born in a litter of 2–4 cubs.{{cite journal |author=Eaton, R.L. |year=1977 |title=Reproductive biology of the leopard |journal=Zoologischer Garten |volume=47 |issue=5 |pages =329–351}} The mortality rate of cubs is estimated at 41–50% during the first year. Predators are the biggest cause for leopard cub mortality during their first year. Male leopards are known to cause infanticide, in order to bring the female back into heat. Intervals between births average 15 to 24 months, but can be shorter, depending on the survival of the cubs.

Females give birth in a cave, crevice among boulders, hollow tree or thicket. Newborn cubs weigh {{Cvt|280-1000|g}}, and are born with closed eyes, which open four to nine days after birth. The fur of the young tends to be longer and thicker than that of adults. Their pelage is also more gray in colour with less defined spots. They begin to eat meat at around nine weeks. Around three months of age, the young begin to follow the mother on hunts. At one year of age, cubs can probably fend for themselves, but will remain with the mother for 18–24 months.{{cite web |url=https://brainmuseum.org/Specimens/carnivora/leopard/index.html |title=Leopard (Panthera pardus); Physical characteristics and distribution |work=Comparative Mammalian Brain Collections |access-date=2008-06-07 |archive-date=2011-01-03 |archive-url=https://web.archive.org/web/20110103183140/http://www.brainmuseum.org/specimens/carnivora/leopard/index.html |url-status=live}} After separating from their mother, sibling cubs may travel together for months. Both male and female leopards typically reach sexual maturity at 2–2⅓ years.

The generation length of the leopard is 9.3 years.{{cite journal |title=Generation length for mammals |last1=Pacifici |first1=M. |last2=Santini |first2=L. |last3=Di Marco |first3=M. |last4=Baisero |first4=D. |name-list-style=amp |last5=Francucci |first5=L. |last6=Grottolo Marasini |first6=G. |last7=Visconti |first7=P. |last8=Rondinini |first8=C. |journal=Nature Conservation |year=2013 |issue=5 |pages=87–94}}

The average life span of a leopard is 12–17 years.

The oldest leopard was a captive female that died at the age of 24 years, 2 months and 13 days.{{cite news |author=Salisbury, S. |date=2014 |url=http://www.palmbeachpost.com/news/news/roxanne-oldest-spotted-leopard-in-captivity-dies-a/ngyYc/ |title=Roxanne, oldest spotted leopard in captivity, dies at Acreage preserve |newspaper=The Palm Beach Post |archive-date=2014-08-11|archive-url=https://web.archive.org/web/20140811165340/http://www.palmbeachpost.com/news/news/roxanne-oldest-spotted-leopard-in-captivity-dies-a/ngyYc/}}

The leopard is listed on CITES Appendix I, and hunting is banned in Botswana and Afghanistan; in 11 sub-Saharan countries, trade is restricted to skins and body parts of 2,560 individuals.

In 2007, a leopard reintroduction programme was initiated in the Russian Caucasus, where captive bred individuals are reared and trained in {{cvt|0.5–0.9|ha}} large enclosures in Sochi National Park; six individuals released into Caucasus Nature Reserve and Alaniya National Park in 2018 survived as of February 2022.{{cite journal |author1=Rozhnov, V.V. |name-list-style=amp |author2=Yachmennikova, A.A. |author3=Dronova, N.A. |author4=Naidenko, S.V. |author5=Hernandez-Blanco, J.A. |author6=Chistopolova, M.D. |author7=Pkhitikov, A.B. |author8=Tembotova, F.A. |author9=Trepet, S.A. |author10=Chestin, I.E. |year=2022 |title=Experience of the leopard re-covering through reintroduction in the Russian Caucasus |journal=Cat News |issue=Special issue |pages=67–71 |url=http://www.catsg.org/fileadmin/filesharing/5.Cat_News/5.3._Special_Issues/5.3.14._SI_15/Cat_News_Special_Issue_15_Persian_leopard_FINAL.pdf#page=67 |access-date=2023-11-11 |archive-date=2023-11-11 |archive-url=https://web.archive.org/web/20231111083835/http://www.catsg.org/fileadmin/filesharing/5.Cat_News/5.3._Special_Issues/5.3.14._SI_15/Cat_News_Special_Issue_15_Persian_leopard_FINAL.pdf#page=67 |url-status=live}}

The leopard is primarily threatened by habitat fragmentation and conversion of forest to agriculturally used land, which lead to a declining natural prey base, human–wildlife conflict with livestock herders and high leopard mortality rates. It is also threatened by trophy hunting and poaching. Contemporary records suggest that the leopard occurs in only 25% of its historical range.{{cite journal|last1=Williams|first1=S. T. |last2=Williams |first2=K. S. |last3=Lewis |first3=B. P. |last4=Hill |first4=R. A. |name-list-style=amp |title=Population dynamics and threats to an apex predator outside protected areas: implications for carnivore management |journal=Royal Society Open Science |date=2017 |volume=4 |issue=4 |pages=161090 |doi=10.1098/rsos.161090 |doi-access=free |pmid=28484625 |pmc=5414262 |bibcode=2017RSOS....461090W}}

Between 2002 and 2012, at least four leopards were estimated to have been poached per week in India for the illegal wildlife trade of its skins and bones.{{Cite report |author1=Raza, R.H. |name-list-style=amp |author2=Chauhan, D.S. |author3=Pasha, M.K.S. |author4=Sinha, S. |date=2012 |title=Illuminating the blind spot: A study on illegal trade in Leopard parts in India (2001–2010) |publisher=TRAFFIC India, WWF India |location=New Delhi |url=https://www.traffic.org/site/assets/files/4130/illuminating_the_blind_spot.pdf |archive-url=https://web.archive.org/web/20200924143713/https://www.traffic.org/site/assets/files/4130/illuminating_the_blind_spot.pdf |archive-date=2020-09-24 |url-status=live}}

In spring 2013, 37 leopard skins were found during a 7-week long market survey in major Moroccan cities.{{cite journal |last1=Bergin |first1=D. |last2=Nijman |first2=V. |name-list-style=amp |year=2014 |title=Open, Unregulated Trade in Wildlife in Morocco's Markets |journal=TRAFFIC Bulletin |volume=26 |issue=1 |pages=65–70 |url=https://www.researchgate.net/publication/267748463 |access-date=2015-03-23 |archive-date=2018-10-31 |archive-url=https://web.archive.org/web/20181031133219/https://www.researchgate.net/publication/267748463 |url-status=live}} In 2014, 43 leopard skins were detected during two surveys in Morocco. Vendors admitted to have imported skins from sub-Saharan Africa.{{Cite journal |last1=Bergin |first1=D. |last2=Nijman |first2=V. |name-list-style=amp |date=2015 |title=Potential benefits of impending Moroccan wildlife trade laws, a case study in carnivore skins |journal=Biodiversity and Conservation |volume=25 |issue=1 |pages=199–201 |doi=10.1007/s10531-015-1042-1 |s2cid=34533018 |url=https://www.researchgate.net/publication/287807462 |access-date=2017-01-11 |archive-date=2020-01-07 |archive-url=https://web.archive.org/web/20200107220433/https://www.researchgate.net/publication/287807462_Potential_benefits_of_impending_Moroccan_wildlife_trade_laws_a_case_study_in_carnivore_skins |url-status=live}}

Surveys in the Central African Republic's Chinko area revealed that the leopard population decreased from 97 individuals in 2012 to 50 individuals in 2017. In this period, transhumant pastoralists from the border area with Sudan moved in the area with their livestock. Rangers confiscated large amounts of poison in the camps of livestock herders who were accompanied by armed merchants. They engaged in poaching large herbivores, sale of bushmeat and trading leopard skins in Am Dafok.{{cite journal |author1=Äbischer, T. |author2=Ibrahim, T. |author3=Hickisch, R. |author4=Furrer, R. D. |name-list-style=amp |author5=Leuenberger, C. |author6=Wegmann, D. |year=2020 |title=Apex predators decline after an influx of pastoralists in former Central African Republic hunting zones |journal=Biological Conservation |volume=241 |page=108326 |doi=10.1016/j.biocon.2019.108326 |bibcode=2020BCons.24108326A |s2cid=213766740 |url=https://doc.rero.ch/record/328409/files/weg_apd.pdf |archive-url=https://web.archive.org/web/20201003124917/https://doc.rero.ch/record/328409/files/weg_apd.pdf |archive-date=2020-10-03 |url-status=live}}

In Java, the leopard is threatened by illegal hunting and trade. Between 2011 and 2019, body parts of 51 Javan leopards were seized including six live individuals, 12 skins, 13 skulls, 20 canines and 22 claws.{{cite journal |author1=Gomez, L. |name-list-style=amp |author2=Shepherd, C.R. |year=2021 |title=The illegal exploitation of the Javan Leopard (Panthera pardus melas) and Sunda Clouded Leopard (Neofelis diardi) in Indonesia |journal=Nature Conservation |volume=43 |issue=43 |pages=25–39 |doi=10.3897/natureconservation.43.59399 |doi-access=free |s2cid=233286106}}

{{Multiple image|direction=vertical|align=right|image1=Sousse museum Smirat-retouched.jpg|caption1=Leopards on the Magerius Mosaic from modern Tunisia |image2=Leopard head hip ornament, Nigeria, Court of Benin, Edo people, late 18th century, bronze, copper, iron - De Young Museum - DSC01037.JPG|upright|caption2=Leopard head ornament from the Court of Benin}}

Leopards have been featured in art, mythology and folklore of many countries. In Greek mythology, it was a symbol of the god Dionysus, who was depicted wearing leopard skin and using leopards as means of transportation. In one myth, the god was captured by pirates but two leopards rescued him.{{cite book |author=Morris, D. |year=2014|title=Leopard|publisher=Reaktion Books|pages=23–24, 31–33, 62, 99, 102, 111 |isbn=9781780233185}} Numerous Roman mosaics from North African sites depict fauna now found only in tropical Africa.{{cite journal |title=The Decline of North Africa Since the Roman Occupation: Climatic or Human? |last1=Murphey |first1=R. |journal=Annals of the Association of American Geographers |volume= XLI |issue=2 |year=1951 |pages=116–132 |doi=10.1080/00045605109352048 |url=http://www.geo.hunter.cuny.edu/courses/geog347/articles/decline_human.pdf |archive-url=https://web.archive.org/web/20060914061431/http://www.geo.hunter.cuny.edu/courses/geog347/articles/decline_human.pdf |archive-date=2006-09-14 |url-status=live}} During the Benin Empire, the leopard was commonly represented on engravings and sculptures and was used to symbolise the power of the king or oba, since the leopard was considered the king of the forest.{{cite web |publisher=British Museum |place=London |title=Benin: an African kingdom |url=https://www.britishmuseum.org/PDF/british_museum_benin_art.pdf |archive-url=https://web.archive.org/web/20080805130342/http://www.britishmuseum.org/PDF/british_museum_benin_art.pdf |archive-date=2008-08-05 |url-status=live |access-date=2016-03-29}} The Ashanti people also used the leopard as a symbol of leadership, and only the king was permitted to have a ceremonial leopard stool. Some African cultures considered the leopard to be a smarter, better hunter than the lion and harder to kill.

In Rudyard Kipling's "How the Leopard Got His Spots", one of his Just So Stories, a leopard with no spots in the Highveld lives with his hunting partner, the Ethiopian. When they set off to the forest, the Ethiopian changed his brown skin, and the leopard painted spots on his skin.{{cite book |last1=Kipling |first1=R. |title=Just So Stories |date=1902 |publisher=Macmillan |chapter=How the Leopard Got His Spots |chapter-url=https://theshortstory.co.uk/devsitegkl/wp-content/uploads/2015/06/Short-stories-Rudyard-Kipling-How-the-Leopard-got-his-Spots.pdf |access-date=2023-05-14 |archive-date=2023-11-29 |archive-url=https://web.archive.org/web/20231129190512/https://theshortstory.co.uk/devsitegkl/wp-content/uploads/2015/06/Short-stories-Rudyard-Kipling-How-the-Leopard-got-his-Spots.pdf |url-status=live}} A leopard played an important role in the 1938 Hollywood film Bringing Up Baby. African chiefs, European queens, Hollywood actors and burlesque dancers wore coats made of leopard skins.

The leopard is a frequently used motif in heraldry, most commonly as passant.{{cite book |author=Haist, M. |title=The Mark of the Beast: The Medieval Bestiary in Art, Life, and Literature |chapter=The Lion, bloodline, and kingship |pages=3–16 |editor=Hassig, D. |publisher=Taylor & Francis |location=London |year=1999 |isbn=978-0-8153-2952-7 |chapter-url=https://books.google.com/books?id=lx09r7EOL_oC&pg=PA10 |access-date=2021-09-07 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505150617/https://books.google.com/books?id=lx09r7EOL_oC&pg=PA10#v=onepage&q&f=false |url-status=live }} The heraldic leopard lacks spots and sports a mane, making it visually almost identical to the heraldic lion, and the two are often used interchangeably. Naturalistic leopard-like depictions appear on the coat of arms of Benin, Malawi, Somalia, the Democratic Republic of the Congo and Gabon, the last of which uses a black panther.{{cite book|title=The International Flag Book in Color |url=https://archive.org/details/internationalfla00pede|url-access=registration|last=Pedersen|first=C. F. |publisher=Morrow |year=1971}}

{{Main article|Leopard attack}}

The Leopard of Rudraprayag killed more than 125 people; the Panar Leopard was thought to have killed over 400 people. Both were shot by British hunter Jim Corbett.{{cite book |title=The Temple Tiger, and More Man-eaters of Kumaon |year=1955 |author=Corbett, J. |author-link=Jim Corbett |publisher=Oxford University Press |place=Oxford}} The spotted devil of Gummalapur killed about 42 people in Karnataka, India.{{cite book |author=Anderson, K. |author-link=Kenneth Anderson (writer) |title=Nine Man-Eaters and one Rogue |publisher=George Allen & Unwin |year=1954 |location=London |chapter=The Spotted Devil of Gummalapur |pages=36–51 |chapter-url=https://archive.org/details/in.ernet.dli.2015.460900/page/n33/mode/2up}}

Image:Female animal trainer and leopard, c1906.jpg

The ancient Romans kept leopards in captivity to be slaughtered in hunts as well as execute criminals. In Benin, leopards were kept and paraded as mascots, totems and sacrifices to deities. Several leopards were kept in a menagerie originally established by King John of England at the Tower of London in the 13th century; around 1235, three of these animals were given to Henry III by Holy Roman Emperor Frederick II.{{cite magazine |last=Owen |first=J. |title=Medieval Lion Skulls Reveal Secrets of Tower of London 'Zoo' |magazine=National Geographic Magazine |date=2005 |url=http://news.nationalgeographic.com/news/2005/11/1103_051103_tower_lions.html |archive-url=https://web.archive.org/web/20051105025912/http://news.nationalgeographic.com/news/2005/11/1103_051103_tower_lions.html |url-status=dead |archive-date=November 5, 2005 |access-date=2007-09-05}} In modern times, leopards have been trained and tamed in circuses.

{{clear}}

• Leopard pattern
• List of largest cats
• Panther (legendary creature)

{{Reflist}}

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• {{cite journal|last1=Zakaria |first1= M. |last2= Sanei|first2=A. |year=2011 |url=https://docs.google.com/viewer?a=v&pid=explorer&srcid=0BzbFC60TvnMcNGExNDJjNzItZmM1MC00ODkzLTkzZWMtNjQ3OTAyMzlmOTcz |title=Conservation and management prospects of the Persian and Malayan leopards|journal=Asia Life Sciences |volume=Supplement 7|pages=1–5 |ref=none}}

{{Commons+cat|Panthera pardus|Panthera_pardus}}

• [http://www.catsg.org/index.php?id=110 IUCN/SSC Cat Specialist Group: Panthera pardus in Africa] and [http://www.catsg.org/index.php?id=557 Panthera pardus in Asia]
• {{Cite EB1911|wstitle=Leopard|short=x}}

{{Carnivora|Fe.}}

{{Mammals in culture}}

{{Taxonbar|from=Q34706|from2=Q47450956}}

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{{DEFAULTSORT:Leopard}}

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